<8) MycoKeys
MycoKeys 121: 111-142 (2025)
DOI: 10.3897/mycokeys.121.157714
Research Article
Four new species of the lichen genus Diorygma (Graphidaceae,
Ostropales) from Guizhou, China
Wei Wu'®, Shu-Hao Jiang”®, Lin-Shan Chai'®, He-Yun Bo'™®, Ruvishika S. Jayawardena*“*®, Shao-Bin Fu™®,
Qing-Feng Meng®®
a fF Ww NY -|
School of Pharmacy, Zunyi Medical University, Zunyi, Guizhou Province 563000, China
College of Agriculture and Biology, Liaocheng University, Liaocheng, Shandong Province 252059, China
Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand
School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand
School of Public Health, Zunyi Medical University, Zunyi, Guizhou Province 563000, China
Corresponding authors: Shao-Bin Fu (fushb@126.com); Qing-Feng Meng (qfmeng@126.com)
OPEN Qaccess
This article is part of:
Exploring the Hidden Fungal Diversity:
Biodiversity, Taxonomy, and Phylogeny of
Saprobic Fungi
Edited by Samantha C. Karunarathna,
Danushka Sandaruwan Tennakoon, Ajay
Kumar Gautam
Academic editor:
Danushka Sandaruwan Tennakoon
Received: 2 May 2025
Accepted: 21 July 2025
Published: 22 August 2025
Citation: Wu W, Jiang S-H, Chai L-S, Bo
H-Y, Jayawardena RS, Fu S-B, Meng
Q-F (2025) Four new species of the
lichen genus Diorygma (Graphidaceae,
Ostropales) from Guizhou, China.
Mycokeys 121: 111-142. https://doi.
org/10.3897/mycokeys.121.157714
Copyright: © Wei Wu et al.
This is an open access article distributed under
terms of the Creative Commons Attribution
License (Attribution 4.0 International - CC BY 4.0).
Abstract
Four new species of the lichen genus Diorygma from China are described based on mor-
phological, chemical, and phylogenetic evidence. Phylogenetic analyses were conduct-
ed using both RAXML and Bayesian posterior probability inference, based on combined
LSU and mtSSU sequences. The characteristic compounds were analyzed by colorimet-
ric reactions and thin-layer chromatography. Diorygma guizhouense is characterized
by small apothecia, a narrow to slightly open disc covered with white pruina, and the
presence of stictic, salazinic, and norstictic acids. Diorygma leigongshanense is char-
acterized by small, oval apothecia, a disc surrounded by entire, raised, and widely open
thalline margins covered with thin, pale yellowish pruina, and the same chemical sub-
stances as D. guizhouense. Diorygma locitonitrus is distinguished by the presence of sal-
azinic acid and hyaline, richly muriform ascospores, notably with distinctly smaller pe-
ripheral cells compared to the central cells. Diorygma weii is characterized by stellately
branched apothecia with a closed to slit-like disc, a milky white thallus tinged with green,
and the presence of only norstictic acid. Detailed morphological descriptions and illus-
trations of the new species are provided, along with a comprehensive species checklist
highlighting the diagnostic characteristics of the known species in this genus.
Key words: 4 new species, lichenized fungi, morphology, muriform, phylogeny
Introduction
Diorygma was described by Eschweiler (1824), who mentioned Opegrapha hi-
eroglyphica as a member of this genus but did not formally designate it as the
type species. Later, Staiger (2002) synonymized Opegrapha hieroglyphica as
Diorygma hieroglyphicum and designated it as the lectotype for Diorygma. The
genus is widely distributed, primarily in tropical regions, and often thrives on
the sheltered or overhanging sides of trees (Staiger 2002; Kalb et al. 2004). It
is characterized by a crustose, white to pale olive-green thallus and lirelliform
to irregularly rounded ascomata with a pruinose disc. The exciple is uncarbon-
111
Wei Wu et al.: Diorygma from China
ized or slightly carbonized, and the hymenium is hyaline, not inspersed, with
branched or anastomosing paraphyses that have a thick gelatinous wall. The
asci are clavate and contain 1-8 spores. The ascospores are hyaline (rarely
brownish), transversely septate to mostly muriform (Li and Jia 2016). Chem-
ically, the genus produces compounds such as norstictic acid, stictic acid, or
the protocetraric acid complex (Kalb et al. 2004; Aptroot et al. 2023).
Kalb et al. (2004) provided a monograph on the genus Diorygma, which in-
cluded 24 recognized species, a detailed taxonomic key, and a phylogenetic tree
based on LSU sequences. Subsequently, Sharma and Makhija (2009a, 2009b)
described four additional species characterized by the presence of norstictic
and salazinic acids as the major secondary metabolites. Later, Sharma and
Khadilkar (2012) reported four more new species from India, two of which
exclusively contain norstictic and salazinic acids. Feuerstein et al. (2014) ex-
panded the diversity of the genus by describing three new species and provid-
ing a revised global key encompassing 52 species. Numerous additional taxa
have been discovered and reported, further increasing diversity (Caceres 2007;
Makhija et al. 2009; Sharma and Makhija 2009a; Lima et al. 2013; Feuerstein et
al. 2014; Sutjaritturakan et al. 2014; Aptroot et al. 2023, 2024). According to the
latest phylogenetic research by Ansil et al. (2023), D. dandeliense B.O. Sharma
& Khadilkar is a synonym of D. karnatakense B.O. Sharma & Khadilkar. Currently,
89 species of the genus Diorygma are recognized worldwide (Table 2). Molecu-
lar data for 15 known species and 4 novel species are available (Table 1).
During a field survey of lichens in Guizhou Province, we discovered several
specimens that formed phylogenetically distinct clades within Diorygma. Fol-
lowing detailed morphological and chemical analyses and comparison with all
known species, we propose these as four new species.
Material and methods
Sample collection and morphological observations
Specimens were collected from the Leigong Shan National Nature Reserve in Leis-
han County and the Yueliangshan Nature Reserve in Congjiang County, Guizhou
Province, China. All voucher specimens are deposited in the Lichen Herbarium of
the Kunming Institute of Botany (KUN-L), Chinese Academy of Sciences, Yunnan,
China. External morphological characteristics of the thallus and ascomata were
examined using a stereomicroscope (OLYMPUS SZX16, Japan) and photographed
with a fitted digital camera (AOR B32, China). Anatomical features, including the
exciple, hymenium, paraphyses, asci, and ascospores, were observed using a light
microscope (OLYMPUS BX53, Japan) based on hand-cut longitudinal sections of
apothecia manually prepared with a razor blade. The sections were immersed in
distilled water, and images were captured with a digital camera (OLYMPUS DP72,
Japan). Lugol's iodine solution (I) was used to stain and examine the hymenium.
Photographic plates were assembled using Adobe Photoshop CC 2019 (Adobe
Systems, USA). Measurements were conducted using ImageJ software (v. 1.50d)
and are presented as (min-—) (x — SD) — (x + SD) (—max), where x is the arithmetic
mean and SD is the standard deviation (rounded to the nearest 0.5 um), followed
by the number of observations (n) when n = 10. The terms ‘min’ and ‘max’ repre-
sent the extreme values observed (Zhurbenko and Diederich 2024).
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714 112
Wei Wu et al.: Diorygma from China
Table 1. Taxon name, voucher/culture, and GenBank accession numbers used in this study. Newly generated sequences
are shown in bold, and “"” indicates holotype strains. Absence of GenBank accession numbers is indicated by “NA”.
Taxon
Diorygma antillarum
D. antillarum
D. antillarum
D. antillarum
D. circumfusu
D. defectoisidiatum
D. defectoisidiatum
D. guizhouense
D. hieroglyphicum
D. junghuhnii
D. junghuhnii
D. junghuhnii
D. junghuhni
D. junghuhnii
D. junghuhnii
D. karnatakense
D. karnatakense
D. karnatakense
D. karnatakense
D. karnatakense
D. leigongshanense
D. locitonitrus
D. microsporum
D. minisporum
D. minisporum
D. aff. minisporum
D. tibellii
D. poitaei
D. pruinosum
D. pruinosum
D. sipmanii
D. tiantaiense
D. toensbergianum
Diorygma sp.
Diorygma sp.
Diorygma sp.
D. weii
Glyphis cicatricosa
G. cicatricosa
Locality
Brazil
USA
El Salvador
El Salvador
Australia
Brazil
Brazil
China
French
Australia
Fiji
Australia
Brazil
India
India
India
India
India
India
India
China
China
USA
Kenya
USA
South Africa
Nicaragua
Nicaragua
Australia
Australia
Costa Rica
China
Brazil
Australia
Fiji
Fiji
China
El Salvador
Kenya
Voucher/Strain
Nelsen 4185 (F)
Nelsen 4037 (F)
Liicking 33018 (F)
Liicking 33019 (F)
Kalb 33922 (Herb. Kalb)
Caceres & Aptroot 28966a’ (ISE)
Caceres & Aptroot 28966b (ISE)
L 0093724" (KUN)
Wirth 26647
Kalb 33937 (Herb. Kalb)
Lumbsch 20539 (F)
Kalb 33931 (Herb. Kalb)
Kalb 33254 (Herb. Kalb)
Mycobiont (MSSRF)
Thallus (MSSRF)
21.26 (AMH)
21.52 (AMH)
21.54 (AMH)
21.55 (AMH)
21.60 (AMH)
L 0093725" (KUN)
L 0093723" (KUN)
Liicking 26504 (F)
Lumbsch 19543v (F)
Liicking 26564 (F)
Medeiros 2106 (DUKE)
Liicking 28533 (F)
Liicking 28538 (F)
Mangold 28g (F)
Kalb 26578 (Herb. Kalb)
Licking 14011 (F)
Jia ZJ19123' (LCUF)
Caceres & Aptroot 42003' (ISE)
Lumbsch 19082I (F)
Lumbsch 205011 (F)
Lumbsch 20513a (F)
L 0093727" (KUN)
Liicking 28047 (F)
Lumbsch 195280 (F)
Chemical component analysis
GenBank accessions number
mtSSU
JX046451
JX046452
JX046453
JX046454
DQ431963
OR270821
OR270822
PQ691396
NA
DQ431962
JX421023
NA
NA
MN944821
MN944822
OP235521
OP235522
OP235523
OP235524
OP235525
PQ847480
PQ847478
JX421024
HQ639598
NA
ON507279
JX421025
HQ639596
NA
NA
DQ431961
NA
OR270820
NA
NA
NA
PQ847479
HQ639610
JX421062
LSU
JX046464
JX046465
NA
JX046467
NA
NA
NA
PQ691395
AY640015
NA
JX421474
AY640017
AY640016
NA
NA
OP235516
OP235517
OP235518
OP235519
OP235520
NA
PQ847477
NA
HQ639626
HQ639665
ON507251
JX421475
HQ639627
JX421476
AY640014
AY640020
MW750692
NA
JX421479
JX421478
JX421477
NA
JX421505
JX421503
The color reactions of the thallus and medulla were tested using the following
reagents: 10% potassium hydroxide (KOH, K), saturated sodium hypochlorite
solution (NaCIO, C), and a saturated solution of p-phenylenediamine in 95%
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714
113
Wei Wu et al.: Diorygma from China
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MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714
Wei Wu et al.: Diorygma from China
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MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714
Wei Wu et al.: Diorygma from China
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MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714
Wei Wu et al.: Diorygma from China
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MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714
Wei Wu et al.: Diorygma from China
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MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714
Wei Wu et al.: Diorygma from China
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MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714
Wei Wu et al.: Diorygma from China
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MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714
Wei Wu et al.: Diorygma from China
ethanol (P). Lichen substances were analyzed by thin-layer chromatography
(TLC) using the solvent C system (formic acid/acetic acid, v/v = 200/30) (Cul-
berson 1972).
DNA extraction, PCR amplification, and sequencing
Genomic DNA was directly extracted from apothecia using a fungal genomic
DNA extraction kit (Solarbio, China), following the manufacturer’s protocol. Mi-
tochondrial small subunit rRNA (mtSSU) sequences were amplified using the
primer pairs mrSSU1/mrSSU3R, while large subunit ribosomal DNA (LSU) se-
quences were amplified using the primer pairs LR5/LROR and AL2R/LR6 (Vil-
galys and Hester 1990; Zoller et al. 1999; Mangold et al. 2008; Kraichak et al.
2015). Polymerase chain reaction (PCR) was performed using a Bio-RAD T-100
thermal cycler in 25 uL reaction volumes, consisting of 12.5 uL of 2x PCR Mix
(including dNTPs mix, Solarbio, China), 8 pL of double-distilled water (ddH,0),
1.0 uL of each 10 mM primer, and 2.5 pL of DNA template. The PCR conditions
were as follows: an initial denaturation at 95 °C for 5 minutes, followed by 38
cycles of denaturation at 94 °C for 45 seconds, annealing at 50 °C for 60 sec-
onds (for mtSSU) or 55 °C for 60 seconds (for LSU), and extension at 72 °C for
90 seconds, with a final extension at 72 °C for 10 minutes and held at 12 °C
(Kraichak et al. 2015). The PCR products were visualized by 1% agarose gel
electrophoresis and subsequently sequenced by Beijing Tsingke Biotech Co.,
Ltd. (Chongqing, China).
Phylogenetic analyses
The sequencing results were evaluated by analyzing chromatograms using
BioEdit Sequence Alignment software (version 7.0.9.0). Forward and reverse
sequences were assembled using ContigExpress software (version 6.0.620.0).
Preliminary taxonomic affiliation and potential sample contamination were
confirmed by BLASTn searches on the NCBI website (https://blast.ncbi.nIm.
nih.gov/Blast.cgi). Newly generated sequences were deposited in GenBank
(Table 1). Additional sequences used for ingroup analysis were retrieved from
the NCBI website (https://www.ncbi.nim.nih.gov), and Glyphis cicatricosa was
selected as the outgroup (Ansil et al. 2023). Phylogenetic analyses were con-
ducted using the OFPT program (Zeng et al. 2023), following its protocol: each
gene region dataset was initially aligned using the ‘auto’ strategy (based on
data size) in MAFFT (Katoh and Standley 2013) and subsequently trimmed us-
ing the ‘gappyout’ method (based on gap distribution) in TrimAl (Capella-Guti-
érrez et al. 2009). Single-gene phylogenetic trees were constructed using the
IQ-TREE Web Server (http://iqtree.cibiv.univie.ac.at/) to confirm well-supported
branches. The best-fit nucleotide substitution models for each dataset were
selected based on the Bayesian Information Criterion (BIC) from 22 common
DNA substitution models with rate heterogeneity, using ModelFinder (Kalyaana-
moorthy et al. 2017). All datasets were concatenated with partition information
for subsequent phylogenetic analyses. Maximum likelihood (ML) analysis was
performed using ultrafast bootstrap approximation (Hoang et al. 2018), com-
bined with the SH-like approximate likelihood ratio test (SH-aLRT) (Guindon et
al. 2010) in IQ-TREE (Nguyen et al. 2015). The consensus tree was summarized
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714 126
Wei Wu et al.: Diorygma from China
based on the extended majority rule. Additionally, ML analysis was conducted
using RAXML-HPC2 on ACCESS (version 8.2.12) in the CIPRES Science Gate-
way (https://www.phylo.org/portal2/login!input.action) with the GTRGAMMA
model and a rapid bootstrap analysis of 1000 replicates (Miller et al. 2010; Sta-
matakis 2014). Bayesian inference was performed using MrBayes (Ronquist
et al. 2012), with two parallel Metropolis-coupled Markov Chain Monte Carlo
runs (one ‘cold’ chain and three heated chains), sampling trees every 1000
generations. The run was automatically terminated when the average standard
deviation of split frequencies dropped below 0.01, and the resulting tree was
summarized after discarding the first 25% of samples as burn-in. The resulting
trees were visualized in FigTree v1.4.4 and further edited in Adobe Illustrator
CC 2019. The new taxon was registered in Index Fungorum (2025) and Faces
of Fungi (Jayasiri et al. 2015).
Result
Phylogenetic analyses
The final dataset comprised 22 taxa and 39 strains/vouchers, with 1562 aligned
characters including gaps (LSU: 760 bp; mtSSU: 802 bp). The RAxML tree was
constructed with a final ML optimization likelihood value of -6039.173664. The
parameters for the GTR+I+G model of combined LSU and mtSSU were as fol-
lows: estimated base frequencies—A = 0.29, C = 0.19, G = 0.27, T = 0.25; substi-
tution rates—AC = 0.66, AG = 2.55, AT = 1.69, CG = 0.76, CT = 7.83, and GT = 1.00.
Bayesian posterior probabilities from MCMC analysis showed a final average
standard deviation of split frequencies of 0.009999. The topologies from both
ML and Bayesian analyses were verified manually and largely concurred (Fig. 1).
Fourteen known species of Diorygma formed a well-supported clade in the
phylogenetic tree. Three new species-—D. leigongshanense, D. locitonitrus, and
D. guizhouense-clustered in a clade with D. tiantaiense Z.F. Jia, with strong
support. In contrast, Diorygma weil was related to D. karnatakense, but this rela-
tionship received low support.
Taxonomy
Diorygma guizhouense Wei Wu & S.B. Fu, sp. nov.
Index Fungorum: IF903744
Facesoffungi Number: FoF 17083
Fig. 2
Etymology. The specific epithet “guizhouense’” refers to the location where the
holotype was collected.
Holotype. KUN-L 0093724
Description. Sexual morph: Thallus corticolous, crustose, thin, tightly at-
tached to the substratum, pale grey to greenish grey, rough, dull, lacking isidia
and soredia, prothallus absent. Apothecia lirelliform, scattered or aggregated,
erumpent, simple or irregularly branched, curved, and either terminally round-
ed or acute, measuring 2-5 mm long and 0.2-0.4 mm wide. Disc narrow to
slightly open, covered with a white pruina. Exciple uncarbonized, brown at
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714 127
Wei Wu et al.: Diorygma from China
Diorygma karnatakense 21. 52 AMH
Diorygma karnatakense 21.55 AMH
100/100! Diorygma karnatakense 21.26 AMH
Diorygma karnatakense 21. 54 AMH
Diorygma karnatakense 21.60 AMH
Diorygma weii L 0093727! KUN
95/0.91-— Diorygma leigongshanense L 0093725" KUN
100/0,29 | Diorygma locitonitrus L 0093723" KUN
2B/L00} =| Diorygma tiantaiense Jia ZJ19123" LCUF
Diorygma guizhouense L 0093724" KUN
Diorygma hieroglyphicum Wirth 26647
Diorygma sp. Lumbsch 205011 F
6 /., Diorygma antillarum Liicking 33018 F
78/-|- Diorygma antillarum Liicking 33019 F
99/100 | Diorygma antillarum Nelsen 4185 F
Diorygma antillarum Nelsen 4037 F
97/0.99 Diorygma defectoisidiatum Caceres & Aptroot 28966a' ISE
9/-| | Diorygma defectoisidiatum Caceres & Aptroot 28966b ISE
Diorygma toensbergianum Caceres & Aptroot 42003' ISE
Diorygma circumfusum Kalb 33922 Herb. Kalb
99/0.92, Diorvema junghuhnii Vhallus MSSRF
Diorygma junghuhnii Mycobiont MSSRF
O4/- Diorygma Junghuhnit Lumbsch 205391 F
99/1 Ook Diorygma junghuhni Kalb 33254 Herb. Kalb
Diorygma junghuhnii Kalb 33931 Herb. Kalb
Diorygma junghuhnii Kalb 33937 Herb. Kalb
Diorygma sp. Lumbsch 20513a F
Diorygma pruinosum Mangold 28g F
99/- Diorygma pruinosum Kalb 26578 Herb. Kalb
Diorygma sp. Lumbsch 190821 F
100/1.00, Diorvema poitaei Licking 28538 F
99/100 Diorygma tibellii Licking 28533 F
100/100 Diorygma sipmanii Liicking 14011 F
Diorygma microsporum Licking 26504 F
97/0.99- Diorygma minisporum Lumbsch 19543v F
100/1.00) | Diorygma aff. minisporum Medeiros 2106 DUKE
Diorygma minisporum Licking 26564 F
100/1.00-— Glyphis cicatricosa Lumbs
Glyphis cicatricosa Liicking
0.02
Figure 1. RAxML analysis based on combined LSU and mtSSU sequence data. Bootstrap support values for maximum
likelihood (ML = 75%) and Bayesian posterior probabilities (PP = 0.90) are shown near the nodes as ML/PP. Glyphis cica-
tricosa (LUcking 28047 F and Lumbsch 195280 F) is used as the outgroup taxon. Newly generated sequences are shown
in bold. Type strains are indicated as '.
apex, pale yellowish brown towards base. Epihymenium brown, 10-30 um
high. Hymenium hyaline, not inspersed, 150-210 um high, I+ weakly blue-vio-
let. Paraphysis anastomosing, filiform, 1-2.5 um wide. Hypothecium weakly
yellowish brown, 15-45 um high. Asci fusiform, 118-222 x 37-82 um, I-.
Ascospores 1/ascus, hyaline, richly muriform, peripheral and central spore
locules of + equal size, ends with gelatinous caps, 24-32 x 5-12 locular,
(104—-)125-119(-214) x (30-)36-58(-77) um (x = 152 x 47 um, n = 20), I-.
Asexual moprh: not observed.
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714 128
Wei Wu et al.: Diorygma from China
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Figure 2. Diorygma guizhouense (KUN-L 0093724, holotype). A-C. Thallus with ascomata; D. Cross section of apothe-
cium; E. Cross section of apothecium (in IKI); H, I. Asci; J-L. Ascospores (in water); F, G. Ascospore (in IKI). Scale bars:
1 mm (A-C); 100 um (D); 200 um (E); 50 um (F); 20 um (G, H-L).
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714
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Wei Wu et al.: Diorygma from China
Chemistry. Thallus K+ reddish brown, C-, KC+ orange, P+ yellow, TLC: stictic
acid, salazinic acid, norstictic acid.
Material examined. CHINA, * Guizhou Province, Congjiang County, Yueliang-
shan Nature Reserve, 25°20'8.56'N, 108°36'19.23"E, 987 m elev., 24 Oct. 2023,
Ze Yang & Bo Liu, Y400 (holotype KUN-L 0093724).
Notes. The new species Diorygma guizhouense is characterized by lirelliform
apothecia, which are erumpent with a narrow to slightly open disc covered by
a white pruina. The exciple is uncarbonized, the hymenium is hyaline, not in-
spersed, and reacts I+ blue-violet. The ascospores, one per ascus, are hyaline,
richly muriform, with peripheral and central spore locules of approximately
equal size, containing 24-32 x 5-12 locular, measuring 125-179 x 36-58 um,
I-. Chemically, this species contains stictic, salazinic, and norstictic acids.
Phylogenetic analysis based on combined LSU and mtSSU sequence data in-
dicates that D. guizhouense is closely related to Diorygma tiantaiense (Fig. 1). A
comparison with D. tiantaiense reveals a 0.97% nucleotide difference in the LSU re-
gion (7/723 bp). Morphologically, D. guizhouense differs by having a narrowly open
disc (vs. fully open) and chemically by having a hymenium that reacts I+ blue-violet
(vs. I-). According to the TLC result, the new species contains stictic and salazinic
acids rather than only norstictic acid in D. tiantaiense (Cui et al. 2024).
According to the taxonomic key provided by Feuerstein et al. (2014),
D. guizhouense is morphologically similar to D. dandeliense, which was later syn-
onymized to D. karnatakense (Ansil et al. 2023). However, D. guizhouense and
D. karnatakense occupy distinct clades in the phylogenetic tree, supporting their
separation at the species level. Chemotaxonomically, TLC analysis shows that
D. guizhouense contains stictic, salazinic, and norstictic acids, whereas D. karna-
takense lacks stictic acid, and the ascospores of D. guizhouense exhibit a negative
iodine reaction (I-), in contrast to the I+ violet reaction reported in related species
(Ansil et al. 2023). The nucleotide comparison reveals clear differences between
the two species: 4.28% (31/725 bp) for LSU and 3.45% (25/725 bp) for mtSSU.
Diorygma leigongshanense Wei Wu & S.B. Fu, sp. nov.
Index Fungorum: IF903745
Facesoffungi Number: FoF 17549
Fig.*3
Etymology. The specific epithet “/eigongshanense’ refers to the location where
the holotype was collected.
Holotype. KUN-L 0093725
Description. Sexual morph: Thallus corticolous, crustose, thin, tightly at-
tached to the substratum, pale grey to greenish grey, rough, dull, lacking isidia
and soredia, prothallus absent. Ascomata lirellate, numerous, oblong to long, +
flexuous, simple or with a few branches, measuring 0.7-2.5 mm long and 0.4-
1.2 mm wide. Dise surrounded by entire raised thalline margins, widely open,
covered with a thin, pale yellowish pruina. Exciple uncarbonized, basally and
laterally brownish. Epihymenium brown, 15-41 um high. Hymenium hyaline, not
inspersed, 160-350 um high, I-. Paraphysis anastomosing, filiform, 1-2.5 um
wide. Hypothecium brown, 20-48 um high. Asci fusiform, 106-202 x 28-58 um,
|-. Ascospores 1/ascus, hyaline, richly muriform, peripheral and central spore
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714 130
Wei Wu et al.: Diorygma from China
Figure 3. Diorygma leigongshanense (KUN-L 0093725, holotype). A-C. Thallus with ascomata; D. Cross section of apoth-
ecium; E-G. Ascus; H-K. Ascospores (in water); L. Ascospore (in IKI). Scale bars: 1 cm (A); 1 mm (B, C); 200 um (D);
20 um (E, F); 50 um (G); 20 um (H-L).
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Wei Wu et al.: Diorygma from China
locules of + equal size, 20-42 x 5-10 locular, (95-)119-170(-194) x (22—)27-
46(-54) um (x = 144 x 37 um, n = 20), |-. Asexual moprh: not observed.
Chemistry. Thallus K+ reddish brown, C-, KC+ yellow, P+ yellow, TLC: stictic
acid, salazinic acid, norstictic acid.
Material examined. CHINA, * Guizhou Province, Leishan County, Leigong
Mountain National Nature Reserve, 26°22'43.16'N, 108°11'42.65'E, 1681 melev.,
on bark, 27 Oct. 2023, Ze Yang & Bo Liu, LGS207 (holotype KUN-L 0093725).
Notes. This species is characterized by its erumpent lirelliform apothecia,
with discs surrounded by entire, raised thalline margins that are widely open
and covered with a thin, pale yellowish pruina. The exciple is uncarbonized, and
the hymenium is hyaline, non-inspersed, and I-. Spores are single per ascus,
hyaline, richly muriform, with peripheral and central spore locules of approx-
imately equal size, 20-42 x 5-10 locular, measuring 119-170 x 27-46 um.
Chemically, this species contains stictic, salazinic, and norstictic acids.
Phylogenetic analysis based on combined LSU and mtSSU sequence data
places D. leigongshanense as closely related to Diorygma tiantaiense (Fig. 1).
However, Diorygma leigongshanense differs chemically by the presence of stic-
tic and salazinic acids (vs. only containing norstictic acid). Morphologically,
D. leigongshanense has lirelliform apothecia, in contrast to the oval, open, and
raised ascocarps of D. tiantaiense (Cui et al. 2024).
Diorygma leigongshanense is morphologically similar to D. rufopruinosum
(A.W. Archer) Kalb, Staiger & Elix. However, it can be distinguished by its asco-
spore septation pattern: in D. leigongshanense, the peripheral cells and central
cells are of similar size, whereas in D. rufopruinosum, the peripheral cells are
noticeably smaller. Diorygma leigongshanense contains stictic, salazinic, and
norstictic acids, while D. rufopruinosum produces protocetraric acid and lacks
stictic acid (Kalb et al. 2004).
Morphologically, Diorygma leigongshanense also shares similarities with
D. chumphonense Sutjaritturakan & K. Kalb, but it can be distinguished by hav-
ing longer ascospores (119-170 um vs. 95-110 um), peripheral cells of equal
size to the central ones (vs. peripheral cells distinctly smaller), lacking stictic
acid (vs. presence), and the I- (vs. I+ blue-violet) (Sutjaritturakan et al. 2014).
Diorygma locitonitrus Wei Wu & S.B. Fu, sp. nov.
Index Fungorum: IF903746
Facesoffungi Number: FoF 17550
Fig. 4
Etymology. “locitonitius” combines “loci,” signifying locality, with “tonitius,” the
Latin word for thunder, to mean “of the locality of thunder,” denoting the loca-
tion where the holotype was found.
Holotype. KUN-L 0093723
Description. Sexual morph: Thallus corticolous, crustose, thin, tightly at-
tached to the substratum, pale grey to greenish grey, rough, dull, lacking isidia
and soredia, prothallus absent. Apothecia lirelliform, scattered or aggregated,
erumpent, simple or irregularly branched, curved, and either terminally rounded
or acute, measuring 0.5-2.5 mm long and 0.2-0.6 mm wide. Dise narrow to
open, covered with a white pruina. Exciple uncarbonized, brown at apex, pale
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714 132
Wei Wu et al.: Diorygma from China
ee
yD .
3 Noa
¥ an ry.
me pt Si ae ag
eh Ss mA a toe
Figure 4. Diorygma locitonitrus (KUN-L 0093723, holotype). A, B. Thallus with ascomata; C. Cross section of apothecium;
D. Cross section of apothecium (in IKI); E, F. Ascus; I-L. Ascospores (in water); G, H. Ascospore (in IKI). Scale bars:
1 mm (A, B); 100 ym (C, D); 50 ym (E, H, |, J); 20 ym (F, J-L).
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714 133
Wei Wu et al.: Diorygma from China
yellowish brown towards base. Epihymenium brown, 10-42 um high. Hymeni-
um hyaline, not inspersed, 180-350 um high, I+ weakly blue-violet. Paraphysis
anastomosing, filiform, 1-2.5 um wide. Asci fusiform, 112-260 x 30-81 um, I-.
Ascospores 1/ascus, hyaline, richly muriform, peripheral cells distinctly smaller
than central ones, 26-40/6-15 locular, (105-)117-189(—247) x (25-)33-60(-
76) um (x = 153 x 47 um, n = 20), I-. Asexual moprh: not observed.
Chemistry. Thallus K+ reddish brown, C-, KC+ orange, P+ yellow, TLC: const-
ictic acid, salazinic acid, norstictic acid.
Material examined. CHINA, * Guizhou Province, Leishan County, Leigong
Mountain National Nature Reserve, 26°22'53.94"N, 108°11'46.76'E, 1771 m elev.,
on bark, 27 Oct. 2023, Ze Yang & Bo Liu, LGS256-1 (holotype KUN-L 0093723).
Notes. This species is characterized by its lirelliform apothecia, which are
erumpent with narrow to open discs covered by a white pruina; the exciple is
uncarbonized, and the hymenium is hyaline, not inspersed, and reacts I+ weakly
blue-violet. The ascospores, one per ascus, are hyaline, richly muriform, with
peripheral cells distinctly smaller than central ones, containing 26-40 x 6-15
locular, measuring 117-189 x 33-60 um, I-. Chemically, this species contains
constictic, salazinic, and norstictic acids.
Phylogenetic analysis based on combined LSU and mtSSU sequence data
places Diorygma locitonitrus as closely related to D. tiantaiense (Fig. 1). Howev-
er, nucleotide comparison of the LSU reveals a difference between D. /ocitoni-
trus of 0.80% (6/758 bp) between the two species. The new taxon is chemically
distinct from the presence of both norstictic and salazinic acids, while D. tian-
taiense contains only norstictic acid. In addition, the hymenium of D. locitoni-
trus exhibits a weakly I+ blue-violet reaction in Lugol's solution, in contrast to
the I- reaction in D. tiantaiense (Cui et al. 2024).
Diorygma locitonitrus is morphologically similar to D. chumphonense but dif-
fers in the larger ascospores (117-189 x 33-60 um vs. 95-110 x 37-40 pm)
and the iodine reaction of the hymenium (I-, vs. + weakly violet). Additionally,
the ascospores of D. /ocitonitrus are |-, in contrast to the I+ violet reaction ob-
served in D. chumphonense (Sutjaritturakan et al. 2014).
Diorygma weii Wei Wu & S.B. Fu, sp. nov.
Index Fungorum: IF903747
Facesoffungi Number: FoF17617
FIG.
Etymology. The species epithet “weii” honors Professor Jiangchun Wei (Chi-
nese Academy of Sciences), a venerable lichenologist, for his pioneering con-
tributions to lichenology in China.
Holotype. KUN-L 0093727
Description. Sexual morph: Thallus corticolous, crustose, thin, tightly at-
tached to the substratum, milky white, with a slight greenish tint, rough, dull,
lacking isidia and soredia, prothallus absent. Apothecia lirelliform, prominent,
stellately branched, curved, and either terminally rounded or acute, measuring
2-7 mm long and 0.1-0.4 mm wide. Dise closed to slit-like, covered with a
thin white pruina. Proper margin conspicuous. Exciple uncarbonized, brown at
apex, pale yellowish brown towards base. Hymenium hyaline, not inspersed,
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714 134
Wei Wu et al.: Diorygma from China
a
pn
So -/ cos
ail
Figure 5. Diorygma weii (KUN-L 0093727, holotype). A-C. Thallus with ascomata; D. Cross section of apothecium; E. Cross
section of apothecium (in IKI); F-H. Ascus; I, J. Ascospores (in water); K. Ascospore (in IKI). Scale bars: 1 mm (A-C);
50 um (D); 100 um (E); 20 um (F-K).
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714
135
Wei Wu et al.: Diorygma from China
150-210 um high, I+ violet. Paraphysis anastomosing, filiform, 1-2 um wide.
Asci fusiform, 60-155 x 18-45 um, I- or I+ violet. Ascospores 1/ascus, hya-
line, richly muriform, peripheral and central spore locules of + equal size, ends
with gelatinous caps, 24—28/6-9 locular, (55—)73-119(-142) x (11-)18-35(-
41) um (x = 96 x 27 um, n = 20), I+ violet. Asexual moprh: not observed.
Chemistry. Thallus K+ reddish brown, C-, KC+ yellow, P+ yellow, TLC: Norst-
ictic acid.
Material examined. CHINA, * Guizhou Province, Leishan County, Leigong Moun-
tain National Nature Reserve, 26°20'33.94'N, 108°17'23.94"E, 831 m elev., on the
bark, 25 Oct. 2023, Ze Yang & Bo Liu, Coll. No. LGS57 (holotype KUN-L 0093727).
Notes. This new species is characterized by its lirelliform apothecia, which
are prominent with a closed-to-slit-like disc covered by a thin white pruina. The
exciple is uncarbonized, and the hymenium is hyaline, not inspersed, and reacts
I+ violet. Spores are single per ascus, hyaline, richly muriform, with peripheral
and central spore locules of approximately equal size, 24-28 x 6-9 locular,
measuring 73-119 x 18-35 um, I+ violet. Chemically, this species contains
only norstictic acid.
Phylogenetic analysis based on the combined data of LSU and mtSSU sequenc-
es indicates that D. weii forms a clade with D. karnatakense (Fig. 1). However,
nucleotide divergence in mtSSU between the two species is 3.52% (23/653 bp).
Morphologically, D. weii has asci containing a single spore, versus 1—8-spored
asci in D. karnatakense. Chemically, D. weii only produces norstictic acid, whereas
D. karnatakense contains both norstictic and salazinic acids (Ansil et al. 2023).
Diorygma weii shares morphological similarities with Diorygma inaequale
and D. dealbatum B.O. Sharma & Makhija. However, both D. inaequale and
D. dealbatum contain both salazinic and norstictic acids, whereas the new spe-
cies produces only norstictic acid (Sharma and Makhija 2009a).
Discussion
Morphological features and chemical compounds are generally used in the
classification of Diorygma species (Kalb et al. 2004). Chemically, most species
exhibit remarkable diversity in secondary metabolites—such as norstictic acid,
stictic acid, cryptostictic acid, and protocetraric acid—which serve as key tax-
onomic markers. Typically, a single species produces one or several of these
characteristic metabolites (Kalb et al. 2004). With the rapid advancement of
molecular technologies, phylogenetic analyses have become an indispensable
tool in species identification and evolutionary studies. Molecular data provide
objective genetic evidence that enables the precise differentiation of morpho-
logically similar or cryptic species, help clarify taxonomic uncertainties includ-
ing synonymies, and facilitate the discovery of new taxa. Phenotypic charac-
teristics alone are sometimes insufficient for resolving taxonomic ambiguities
due to environmental influences and developmental stage. In contrast, genetic
data are relatively stable. DNA sequences from LSU and mtSSU are usually
considered reliable molecular markers, and phylogenetic trees based on these
genes can confirm the monophyletic nature of species defined by morphology
and correct misclassifications caused by overlapping features of sporangia or
ascospores within Diorygma (Cui et al. 2024; Aptroot et al. 2023). Therefore,
this study adopts an integrative taxonomic approach, combining phylogenet-
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714 136
Wei Wu et al.: Diorygma from China
ic analyses, morphological characteristics, and chemical profiling to achieve a
comprehensive and accurate classification of new Diorygma species.
Most Diorygma species have stable phylogenetic positions with stronger sta-
tistical support in multigene datasets than in single-gene analyses (Ansil et al.
2023). The phylogenetic tree based on LSU + mtSSU sequence data provided
high-resolution species delimitation within Diorygma. Diorygma guizhouense,
D. leigongshanense, and D. locitonitrus formed a highly supported clade (ML/
PP = 90/0.90) with D. tiantaiense. Although D. weii and D. karnatakense formed
a distinct clade, this relationship received weak support (ML/PP < 70/0.90). All
four proposed species occupy distinct phylogenetic positions.
Approximately 90 Diorygma species are currently recognized and categorized
by ascospore number into two morphological groups (Table 2). Among the 38
monosporate species, our four new taxa all belong to this group. Thirteen mono-
sporate species produce salazinic acid—a feature shared by D. guizhouense and
D. leigongshanense. These two species are further distinguished by co-occurring
salazinic and stictic acids, a combination previously documented only in D. an-
gusticarpum Sutjaritturakan & Kalb and D. salazinicum Sutjaritturakan & Kalb.
Morphological and chemical comparisons reveal that D. guizhouense can
be distinguished from D. angusticarpum by its larger ascospores (125-179 x
36-58 um vs. 90-110 x 30-37 um) and the presence of norstictic acid (vs.
cryptostictic acid) (Sutjaritturakan et al. 2014). Additionally, D. guizhouense dif-
fers from D. salazinicum in having a narrower disc (vs. widely opened discs), a
weakly I+ blue-violet hymenium reaction (vs. I-), and the presence of norstictic
acid (vs. cryptostictic acid) (Sutjaritturakan et al. 2014).
Comparative analysis reveals that D. leigongshanense exhibits unique diag-
nostic features when contrasted with similar species. Compared to D. angus-
ticarpum, the new species displays markedly larger ascospores (119-170 x
27-46 um vs. 90-110 x 30-37 um), a distinctly broader disc (vs. slit-like), an
I- hymenium (vs. I+ violet), and contains norstictic acid (vs. cryptostictic acid)
(Sutjaritturakan et al. 2014). These chemical and morphological differences
also distinguish D. leigongshanense from D. salazinicum, particularly in the io-
dine reaction patterns of the ascospores (I- vs. I+ violet) and secondary metab-
olite profiles (norstictic acid vs. cryptostictic acid) (Sutjaritturakan et al. 2014).
Taxonomic evaluation of ascospore septation (cell size) in monosporate
Diorygma species revealed two distinct morphological types: those with pe-
ripheral cells smaller than central cells and those with cells of equal size. The
newly described D. locitonitrus possesses the former pattern, a characteristic
shared with nine other monosporate species. Notably, only three previously
documented species—D. reniforme (Fée) Kalb, Staiger & Elix, D. rufopruinosum,
and D. salvadoriense Kalb, Staiger & Elix—exhibit both this septation pattern
and the presence of salazinic acid. Diorygma locitonitrus can be distinguished
from these chemically similar species by the absence of protocetraric acid (vs.
present in D. reniforme and D. rufopruinosum) and the absence of pycnidia (vs.
present in D. salvadoriense) (Kalb et al. 2004). These consistent differences in
secondary metabolite profiles and reproductive structures provide robust crite-
ria for delimiting D. locitonitrus from related taxa.
Among monosporate Diorygma species, seven taxa—D. australasicum (Elix)
Licking, Elix & A.W. Archer, D. isabellinum (Zahlbr.) Z.F. Jia & Liicking, D. longil-
irellatum B.O. Sharma & Makhija, D. roseopruinatum Papong, Lucking & Parn-
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714 137
Wei Wu et al.: Diorygma from China
men, D. soozanum (Zahlbr.) M. Nakan. & Kashiw., D. spilotum (Stirt.) Pushpi
Singh & Kr.P. Singh, and D. tiantaiense—exclusively produce norstictic acid (Ta-
ble 2), warranting detailed comparison with D. weii, which shares this chemical
profile. Diorygma weii is distinguished by its unique thallus morphology (milky
white with a slight greenish tint) and apothecial features (stellately branched
with closed to slit-like discs). Key diagnostic characters that distinguish D. weii
from related species include: (1) From D. roseopruinatum by thallus color (milky
white vs. light grey), and disc exposure and pruinosity (closed, slit-like white
pruinose vs. narrow brown with pink-red pruina) (Papong et al. 2014); (2) From
D. australasicum by the absence of isidia (Archer and Elix 2009); (3) From D.
spilotum by branching of lirellae (prominent stellately branched vs. simple im-
mersed) and disc pruinosity (white pruinose vs. epruinose) (Singh and Singh
2017); (4) From D. isabellinum by branching of lirellae (stellately vs. single/rare-
ly branched) and iodine reaction (I+ violet hymenium vs. I-) (Jia and Liicking
2017); (5) From D. longilirellatum by thallus color (milky white vs. greenish-grey)
and disc exposure (closed vs. slightly open) (Sharma and Makhija 2009b); (6)
From D. tiantaiense by smaller ascospores (73-119 x 18-35 um vs. 120-210 x
35-60 um), disc exposure (closed vs. open), and iodine reactions (I+ violet vs.
I- for both hymenium and ascospores) (Cui et al. 2024); (7) From D. soozanum
by narrower ascospores (73-119 x 18-35 um vs. 110-140 x 35-45 um) and
disc exposure (closed vs. initially narrow then wide) (Kalb et al. 2004).
This study describes four new species of the lichen genus Diorygma from Chi-
na. Each species exhibits distinctive morphological features and secondary me-
tabolite profiles, confirmed through colorimetric tests and thin-layer chromatog-
raphy. These findings contribute to a deeper understanding of Diorygma diversity
in China and underscore the value of combining traditional taxonomy with mo-
lecular and chemotaxonomic approaches. A comprehensive species checklist is
also provided to aid in the identification and comparison of known Diorygma taxa.
Additional information
Conflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Use of Al
No use of Al was reported.
Funding
No funding was reported.
Author contributions
Wei Wu and Shao-Bin Fu designed the experiments and structured the manuscript. Wei
Wu conducted the experiments, analyzed the data, and drafted the manuscript. He-Yun
Bo and Lin-Shan Chai conducted part of the molecular and chemical experiments. Qing-
Feng Meng, Shu-Hao Jiang, Ruvishika S. Jayawardena, and Shao-Bin Fu contributed to
data analysis and manuscript revision.
MycoKeys 121: 111-142 (2025), DOI: 10.3897/mycokeys.121.157714 138
Wei Wu et al.: Diorygma from China
Author ORCIDs
Wei Wu © https://orcid.org/0009-0009-1 191-6667
Shu-Hao Jiang ® hitps://orcid.org/0009-0009-8619-2667
Lin-Shan Chai © https://orcid.org/0009-0005-2458-0454
He-Yun Bo © https://orcid.org/0009-0003-8641-1619
Ruvishika S. Jayawardena © https://orcid.org/0000-0001-7702-4885
Shao-Bin Fu © https://orcid.org/0000-0001-9932-1346
Qing-Feng Meng ® https://orcid.org/0000-0001-9814-8238
Data availability
All of the data that support the findings of this study are available in the main text.
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